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Nomenclature and Distribution of the Species of the Porcupinefish Family

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Nomenclature and Distribution of the Species of the Porcupinefish Family Memoirs of Museum Victoria 63(1): 77–90 (2006) ISSN 1447-2546 (Print) 1447-2554 (On-line) http://www.museum.vic.gov.au/memoirs/index.asp Nomenclature and distribution of the species of the porcupinefi sh family Diodontidae (Pisces, Teleostei) JEFFREY M. LEIS Ichthyology, Australian Museum, 6 College St, Sydney, NSW 2010, Australia (e-mail: jeffl @austmus.gov.au) Abstract Leis, J.M. 2006. Nomenclature and distribution of the species of the porcupinefi sh family Diodontidae (Pisces, Teleostei). Memoirs of Museum Victoria 63(1): 77–90. The tetraodontiform fi sh family Diodontidae is widely distributed in tropical and temperate marine waters. The family has more than 70 nominal species, over 60% of which were described in the 100 years following Linnaeus. As a consequence, many descriptions are less than detailed, and many types are no longer extant, if they existed at all. The high incidence of synonymy, the many ʻoldʼ descriptions and the wide geographical distributions of the species has led to a great deal of confusion. The present study, based on examination of diodontid holdings in 29 major collections, and including the extant types of all but two of the nominal species, attempts to clarify the nomenclature and distribution of the species of the family. Although some species boundaries are not entirely clear, only 18 or 19 of the nominal species are herein regarded as valid (one as a subspecies). Tentative assignments of species to genera maintain current usage. Final assignments to genera must await a cladistic analysis of relationships within the family. Four species are circumtropical, four species (plus a subspecies) are confi ned to the Atlantic and appear to form a species group, four species are widely distributed in the tropical Indo-Pacifi c, two species are confi ned to tropical Australasia, and three are endemic to temperate Australia. One species described from New Zealand either occurs also in Australia, or is a synonym of an Australian species. Synonymies, a key to the recognized species and a table of the identities of nominal species are provided. Keywords Tetraodontiformes, burrfi sh, Allomycterus, Chilomycterus, Cyclichthys, Dicotylichthys, Diodon, Lophodiodon, Tragulichthys Introduction This paper lists the senior synonym of each morphologically- defi ned species, followed by the junior synonyms. Brief The porcupinefi sh family Diodontidae contains about justifi cations for synonymies are provided, as are descriptions 19 species in seven or eight genera of warm to temperate seas. of the distribution of the species based on material examined There are about 75 nominal species in the family, and several or on identifi able literature records. In addition to the key of the species have very wide distributions. The species are provided here, regional keys to the species, and illustrations of conspicuous, readily captured, of unusual morphology, and them, can be found in Leis (1986, ref. 5686 – western Indian have been the focus of interest by naturalists from ancient Ocean); Leis (2001, ref. 26318 – western central Pacifi c Ocean: times. Several species have pelagic stages that reach large this key also covers all species in the eastern Indian Ocean); sizes and differ in appearance from the demersal adults. Allen and Robertson (1994, ref. 22193 – eastern Pacifi c Ocean); All this has led to a great deal of confusion as to the number Leis (2003, ref. 27121 – western central Atlantic Ocean); Leis of species, their distributions and the correct names for (in press, eastern central Atlantic Ocean). The key in Leis them. The purpose of this paper is to clarify a number of (2001, ref. 26318) includes all Indo-Pacifi c species and genera nomenclatural issues and the distributions of the species. recognized herein, except the two temperate Australasian Due to their wide distributions, the species are obvious targets species Diodon nicthemerus Cuvier and Allomycterus pilatus for molecular genetic studies, and it is reasonable to expect Whitley (for these, see Kuiter, 1993, ref. 23929, or Gomon et that some taxa that are currently, on the basis of morphology al., 1994, ref. 22532 ). alone, considered to represent a single, widespread species will be subdivided once genetic studies are undertaken. Materials and methods Conversely, some Atlantic taxa have less than clear separations and may eventually be considered conspecifi c. Therefore, Abbreviations of fi ns are as follows: D, dorsal; A, anal; P, it is important to lay some groundwork for these expected pectoral; C, caudal. The spines mentioned are the dermal spines future studies. (i.e., modifi ed scales): fi ns of diodontids lack spines. These 78 Jeffrey M Leis dermal spines have subdermal bases (or roots) that have either Pacifi c genera, three of which are confi ned to Australasia, two approximately opposing bases upon which the exposed contain species that have a mixture of fi xed and erectile spines spine pivots when it elevates, or three (occasionally four) – Allomycterus, Dicotylichthys, Lophodiodon and Tragulichthys broadly more or less equidistantly spaced bases that render the – are usually recognized (see Gomon, 1994, ref. 22532 ; Leis, exposed spine immobile. The exposed portion of the spine 2001, ref. 26318) and are in this paper. Chilomycterus sensu lato varies in length and shape, but erectile spines are generally (about ten species with nearly all dermal spines fi xed and round in cross-section, whereas fi xed spines can vary from immovable) has been more problematical. Tyler (1980, ref. round to compressed in cross-section. See Leis (1978, ref. 5529; 4477) recognized three groupings of Chilomycterus: 1) ʻAtlantic 1986, ref. 5686; or 2001, ref. 26318) for more information on Chilomycterusʼ (fi ve species confi ned to the Atlantic, and called spine morphology. Behind the massive beak-like jaws of by him antennatus, antillarum, mauretanicus, schoepfi i and diodontids is a grinding, or trituration, plate formed by the spinosus); 2) what I call herein ʻCircumtropical Chilomycterusʼ fused premaxillae and dentaries. This plate is often armed with (a circumtropical group considered by Tyler to consist of four transverse plates of teeth, called trituration teeth. species called by him affi nis, atinga, reticulatus and tigrinus); Specifi c information on types is included only if it and 3) ʻIndo-Pacifi c Chilomycterusʼ (considered by Tyler to supplements or corrects information in Eschmeyer (2005; 17 consist only of orbicularis). I agree with Tyler (1980, ref. 4477) Oct 2005 version). To keep the literature cited list to a manageable that nominal species in each of these three groups are length, it includes only references not included in Eschmeyerʼs morphologically more similar to each other than they are to (2005) on-line database (http://www.calacademy.org/research/ species in the other groups. The type species of Chilomycterus ichthyology/catalog/fi shcatmain.asp). The Eschmeyer reference is Diodon reticulatus Linnaeus (1758, ref. 2787); thus, if these number is included with the text citation. Information on Diodon groupings prove to be valid at the generic level, the circumtropical is included in Leis (1978, ref. 5529) and in Leis and Bauchot group becomes Chilomycterus, and I use it in that sense herein. (1984, ref. 12539). For Diodon, only information on species Cyclichthys typically is used for several Indo-Pacifi c species described since 1978 and information updating species (including two not mentioned by Tyler [1980, ref. 4477]), and distributions is included here. the type of Cyclichthys is the Indo-Pacifi c orbicularis; thus, the I examined specimens of diodontids from the following Indo-Pacifi c grouping can be considered Cyclichthys for the institutions (codes after Leviton et al., 1985, ref. 9683): AMS, purposes of this paper. The Atlantic group of species, regarded ANSP (loans based on holdings list), BMNH, BPBM, CAS, by Tyler to be the most phylogenetically basal, is nearly always CSIRO, FAKU, FMNH (loans based on holdings list), FRSKU, included in Chilomycterus. If these Atlantic species were LACM, MCZ, MNHN, NMNZ, NMV, NSMT, NTM, QM, removed from Chilomycterus, the generic name available for RMNH, ROM, RUSI, SAMA, SIO, SMF, UA, USNM, WAM, them is Lyosphaera, based on the unique pelagic stage found in ZMA, ZMB, ZMUC. Distributions are based primarily on at least some members of this group. Unfortunately, the identity museum specimens examined, but are supplemented with of the type species is not clear (on the basis of distribution, reliable literature accounts. schoepfi i seems most likely). Lyosphaera has not been used as a generic name for these fi ve Atlantic species, and until a full Results cladistic analysis is performed on the group, its use is not recommended. For the purposes of this paper, I use Tylerʼs term, Family Diodontidae “Atlantic Chilomycterus”, to identify this grouping. Diagnosis. Small to medium-sized fi shes to 1 m in length, commonly 20–50 cm. Body wide and capable of great infl ation, Chilomycterus (ex Bibron) Brisout de Barneville, 1846 (sensu covered with massive spines that may be quite long; spines with stricto) large bases, or roots, under the skin; long spines usually erectile Chilomycterus (ex Bibron) Brisout de Barneville, 1846 (type and two-rooted, short spines usually fi xed in erect position by species, Diodon reticulatus Linnaeus) their three-rooted bases. Head broad and blunt; gill opening a Cyanichthys Kaup, 1855 (type species is D. coeruleus [non-D. relatively small, vertical slit immediately before pectoral-fi n caeruleus Quoy and Gaimard] Kaup = D. reticulatus Linnaeus, 1758) base; nasal organ usually in small tubes located in front of Diagnosis. All spines fi xed, with long subdermal bases but large eyes; mouth large, wide and terminal; teeth fused to form short or absent external spines (relatively smaller in larger a strong, beak-like crushing structure without a median suture individuals); some spines on top of head with 4 bases; 10 C dividing the upper and lower jaws into left and right halves.
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